土壤酸化对景天三七叶片光合特性及超微结构的影响
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摘要
以景天三七(Sedum aizoon L.)为材料,通过浓硫酸模拟土壤酸化,研究了不同土壤酸化处理(pH 3.4、4.6、5.5)对景天三七叶片光合特性及超微结构的影响。结果表明,随着土壤酸化加剧,土壤中铝离子溶出量迅速增加,叶片总叶绿素含量则逐渐降低,而细胞相对电导率和丙二醛含量逐渐增高。在轻度(pH 5.5)和中度(pH 4.6)酸化下植株光合及荧光参数与CK(pH 6.6)相比无显著差异,叶绿体超微结构完整;在重度(pH 3.4)酸化下,光合及荧光参数出现明显阈值响应,P_n、C_i、T_r、G_s、F_o、F_m、Φ_(PSⅡ)、qP、F_v/F_m迅速降低,NPQ、L_s则显著增高,叶绿体不再沿细胞膜均匀分布,并在细胞质中扎堆聚集,基粒片层出现明显间隙,部分类囊体膜模糊不清。结果还表明,土壤酸化促进了叶片中游离脯氨酸、可溶性蛋白,以及SOD、CAT、GR和APX活性的增加,可溶性糖则呈现先增加后下降趋势。由此可见,土壤酸化导致植株叶片膜质过氧化的原因可能是促进了土壤中铝离子的析出;轻中度土壤酸化并未引起叶片光合及超微结构的变化,重度土壤酸化则会导致叶绿体结构和功能遭受破坏,光合能力降低;同时景天三七可通过诱导渗透物质的增加和抗氧化酶活性的增强以防御土壤酸化的伤害。
In this study, the effect of soil acid stress on photosynthetic characteristics and ultrastructure of Sedum aizoon L. leaves were studied by simulating soil acidification under different soil pH values(3.4, 4.6, 5.5). The results showed that with the intensification of soil acidification, the dissolution of aluminum ions in soil increased rapidly, while the total chlorophyll content in leaves decreased gradually and the relative cell conductivity and malondialdehyde content increased gradually. Under mild(pH 5.5) and moderate(pH 4.6) acidification, plant photosynthetic and fluorescence parameters were not significantly different from CK(pH 6.6), and chloroplast ultrastructure was intact. Under severe(pH 3.4) acidification, photosynthesis and fluorescence parameters showed obvious threshold response, P_n, C_i, T_r, G_s, F_o, F_m, Φ_(PSⅡ), qP, F_v/F_m fell rapidly, NPQ and L_s were significantly higher, and the chloroplast was no longer uniformly distributed along the cell membrane, and the cluster aggregated in the cytoplasm, the granum lamella appeared obvious gap, parts of the internal capsule membrane blurred. Thus, the reason of soil acidification leading to membranous peroxidation of plant leaves might be that it promoted the increase of aluminum ions in soil. Mild and moderate soil acidification did not cause changes in leaf photosynthesis and ultrastructure, while severe soil acidification would lead to destruction of chloroplast structure and function and decrease of photosynthetic capacity. At the same time, the plant can defend against the harm of soil acidification by inducing the increase of osmotic substances and the enhancement of antioxidant enzyme activity.
In this study, the effect of soil acid stress on photosynthetic characteristics and ultrastructure of Sedum aizoon L. leaves were studied by simulating soil acidification under different soil pH values(3.4, 4.6, 5.5). The results showed that with the intensification of soil acidification, the dissolution of aluminum ions in soil increased rapidly, while the total chlorophyll content in leaves decreased gradually and the relative cell conductivity and malondialdehyde content increased gradually. Under mild(pH 5.5) and moderate(pH 4.6) acidification, plant photosynthetic and fluorescence parameters were not significantly different from CK(pH 6.6), and chloroplast ultrastructure was intact. Under severe(pH 3.4) acidification, photosynthesis and fluorescence parameters showed obvious threshold response, P_n, C_i, T_r, G_s, F_o, F_m, Φ_(PSⅡ), qP, F_v/F_m fell rapidly, NPQ and L_s were significantly higher, and the chloroplast was no longer uniformly distributed along the cell membrane, and the cluster aggregated in the cytoplasm, the granum lamella appeared obvious gap, parts of the internal capsule membrane blurred. Thus, the reason of soil acidification leading to membranous peroxidation of plant leaves might be that it promoted the increase of aluminum ions in soil. Mild and moderate soil acidification did not cause changes in leaf photosynthesis and ultrastructure, while severe soil acidification would lead to destruction of chloroplast structure and function and decrease of photosynthetic capacity. At the same time, the plant can defend against the harm of soil acidification by inducing the increase of osmotic substances and the enhancement of antioxidant enzyme activity.
引文
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[18] 熊庆娥.植物生理学实验教程[M].成都:四川科学技术出版社,2013.
[19] 明华,胡春胜,张玉铭,等.浸提法测定玉米叶绿素含量的改进[J].玉米科学,2007,15(4):93-95.MING H,HU C S,ZHANG Y M,et al.Improved extraction methods of chlorophyll from maize[J].Journal of Maize Sciences,2007,15(4):93-95.(in Chinese with English abstract)
[20] 李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[21] 李忠光,李江鸿,杜朝昆,等.在单一提取系统中同时测定五种植物抗氧化酶[J].云南师范大学学报(自然科学版),2002,22(6):44-48.LI Z G,LI J H,DU C K,et al.Simultaneous measurement of five antioxidant enzyme activities using a single extraction system[J].Journal of Yunan Normal University (Natural Sciences Edition),2002,22(6):44-48.(in Chinese with English abstract)
[22] NAKANO Y,ASADA K.Hydrogen peroxide is scavenged by ascorbate specific peroxidase in spinach chloroplasts[J].Plant and Cell Physiology,1981,22(5):867-880.
[23] MA F W,CHENG L L.The sun-exposed peel of apple fruit has higher xanthophyll cycle-dependent thermal dissipation and antioxidants of the ascorbate-glutathione pathway than the shade peel[J].Plant Science,2003,165(4):819-827.
[24] BERRY J A,DOWNTON W J S.Environmental regulation of photosynthesis[M]//GOVINDJEE.Photosynthesis.New York:Academic Press,1982:263-342.
[25] CUDDY W S,SUMMERELL B A,GEHRINGER M M,et al.Nostoc,microcoleus and leptolyngbya inoculums are detrimental to the growth of wheat (Triticum aestivum L.) under salt stress[J].Plant and Soil,2013,370(1/2):317-332.
[26] ROSENBERG M B,BUTCHER D J.Investigation of acid deposition effects on southern appalachian red spruce (Picea rubens) by determination of calcium,magnesium,and aluminum in foliage and surrounding soil using ICP-OES[J].Instrumentation Science and Technology,2010,38(5):341-358.
[27] LI Z,PENG Y,MA X.Different response on drought tolerance and post-drought recovery between the small-leafed and the large-leafed white clover (Trifolium repens L.) associated with antioxidative enzyme protection and lignin metabolism[J].Acta Physiologiae Plantarum,2013,35(1):213-222.
[28] MORGAN J M,HARE R A,FLETCHER R J.Genetic variation in osmoregulation in bread and durum wheats and its relationship to grain yield in a range of field environments[J].Australian Journal of Agricultural Research,1986,37(5):411-420.
[29] 井大炜,邢尚军,杜振宇,等.干旱胁迫对杨树幼苗生长、光合特性及活性氧代谢的影响[J].应用生态学报,2013,24(7):1809-1816.JING D W,XING S J,DU Z Y,et al.Effects of drought stress on the growth,photosynthetic characteristics,and active oxygen metabolism of poplar seedlings[J].Chinese Journal of Applied Ecology,2013,24(7):1809-1816.(in Chinese with English abstract)
[2] BINKLEY D,DRISCOLL C T,ALLEN H I,et al.Acidic deposition and forest soils[M].New York:Springer-Verlag,1989:1-12,65-85.
[3] CREGAN P D,SCOTT B J.Soil acidification-an agricultural and environmental problem[M]//PRATLEY J E,ROBERTSON A.Agriculture and the environmental imperative.Melbourne:CSIRO Publishing,1998:98-128.
[4] GUO J H,LIU X J,ZHANG Y,et al.Significant acidification in major Chinese croplands[J].Science,2010,327 (5968):1008-1010.
[5] 韩江培.设施栽培条件下土壤酸化与盐渍化耦合发生机理研究[D].杭州:浙江大学,2015.HAN J P.The mechanisms for the coexistence of soil acidification and salinization under greenhouse cultivation[D].Hangzhou:Zhejiang University,2015.(in Chinese with English abstract)
[6] LU S G,BAI S Q,XUE Q F.Magnetic properties as indicators of heavy metals pollution in urban top soil:a case study from the city of Luoyang,China[J].Geophysical Journal International,2007,171(2):568-580.
[7] 罗敏.江苏省茶园土壤酸化现状及其影响因素研究[D].南京:南京农业大学,2006.LUO M.Study on soil acidification status and influence factors of tea plantations in Jiangsu Province[D].Nanjing:Nanjing Agricultural University,2006.(in Chinese with English abstract)
[8] TAYLOR G J.Current views of the aluminum stress response:the physiological basis of tolerance[J].Plant Biochemistry and Physiology,1991,10:57-95.
[9] SCHWARZEROVá K,ZELENKOVá S,NICK P,et al.Aluminuminduced rapid changes in the microtubular cytoskeleton of tobacco cell line[J].Plant & Cell Physiology,2002,43:207-216.
[10] 黄丽媛.油茶对铝的吸收积累及其耐性机理研究[D].长沙:中南林业科技大学,2017.HUANG L Y.Mechanisms of aluminum uptake and tolerance in Camellia oleifera Abel.[D].Changsha:South University of Forestry and Technology,2017.(in Chinese with English abstract)
[11] 应小芳.大豆耐铝毒的营养和生理机理研究[D].金华:浙江师范大学,2005.YING X F.Studies on nutritional and physiological mechanisms of soybean as affected with aluminum[D].Jinhua:Zhejiang Normal University ,2005.(in Chinese with English abstract)
[12] 易杰祥,吕亮雪,刘国道.土壤酸化和酸性土壤改良研究[J].华南热带农业大学学报,2006,12(1):23-28.YI J X,LYU L X,LIU G D.Research on soil acidification and acidic soil's melioration[J].Journal of South China University of Tropical Agriculture,2006,12(1):23-28.(in Chinese with English abstract)
[13] 王嵩,马杰,赵维,等.黔西北景天三七高产栽培优化研究[J].北方园艺,2014 (11):147-149.WANG S,MA J,ZHAO W,et al.Research on optimazation growing cutivation of Sedum aizoon L.in Northwest Guizhou[J].Northern Horticulture,2014(11):147-149.(in Chinese with English abstract)
[14] 刘晔峰,马峥嵘,吴晓琴,等.养心草的有效成分及功能性化合物研究进展[J].中华中医药杂志,2012,27(8):2135-2138.LIU Y F,MA Z R,WU X Q,et al.Research progress of active principle and functional compounds of Sedum aizoon L.[J].China Journal of Traditional Chinese Medicine and Pharmacy,2012,27(8):2135-2138.(in Chinese with English abstract)
[15] 董庆武.药食蔬菜养心菜的栽培技术[J].现代农业,2006(11):27.DONG Q W.The cultivation techniques of vegetable and vegetable cultivation[J].Modern Agriculture,2006(11):27.(in Chinese)
[16] 鲁如坤.土壤农业化学分析方法[M].北京:中国农业科技出版社,2000:12-14.
[17] 王学奎.植物生理生化实验原理和技术[M].北京:高等教育出版社,2006.
[18] 熊庆娥.植物生理学实验教程[M].成都:四川科学技术出版社,2013.
[19] 明华,胡春胜,张玉铭,等.浸提法测定玉米叶绿素含量的改进[J].玉米科学,2007,15(4):93-95.MING H,HU C S,ZHANG Y M,et al.Improved extraction methods of chlorophyll from maize[J].Journal of Maize Sciences,2007,15(4):93-95.(in Chinese with English abstract)
[20] 李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[21] 李忠光,李江鸿,杜朝昆,等.在单一提取系统中同时测定五种植物抗氧化酶[J].云南师范大学学报(自然科学版),2002,22(6):44-48.LI Z G,LI J H,DU C K,et al.Simultaneous measurement of five antioxidant enzyme activities using a single extraction system[J].Journal of Yunan Normal University (Natural Sciences Edition),2002,22(6):44-48.(in Chinese with English abstract)
[22] NAKANO Y,ASADA K.Hydrogen peroxide is scavenged by ascorbate specific peroxidase in spinach chloroplasts[J].Plant and Cell Physiology,1981,22(5):867-880.
[23] MA F W,CHENG L L.The sun-exposed peel of apple fruit has higher xanthophyll cycle-dependent thermal dissipation and antioxidants of the ascorbate-glutathione pathway than the shade peel[J].Plant Science,2003,165(4):819-827.
[24] BERRY J A,DOWNTON W J S.Environmental regulation of photosynthesis[M]//GOVINDJEE.Photosynthesis.New York:Academic Press,1982:263-342.
[25] CUDDY W S,SUMMERELL B A,GEHRINGER M M,et al.Nostoc,microcoleus and leptolyngbya inoculums are detrimental to the growth of wheat (Triticum aestivum L.) under salt stress[J].Plant and Soil,2013,370(1/2):317-332.
[26] ROSENBERG M B,BUTCHER D J.Investigation of acid deposition effects on southern appalachian red spruce (Picea rubens) by determination of calcium,magnesium,and aluminum in foliage and surrounding soil using ICP-OES[J].Instrumentation Science and Technology,2010,38(5):341-358.
[27] LI Z,PENG Y,MA X.Different response on drought tolerance and post-drought recovery between the small-leafed and the large-leafed white clover (Trifolium repens L.) associated with antioxidative enzyme protection and lignin metabolism[J].Acta Physiologiae Plantarum,2013,35(1):213-222.
[28] MORGAN J M,HARE R A,FLETCHER R J.Genetic variation in osmoregulation in bread and durum wheats and its relationship to grain yield in a range of field environments[J].Australian Journal of Agricultural Research,1986,37(5):411-420.
[29] 井大炜,邢尚军,杜振宇,等.干旱胁迫对杨树幼苗生长、光合特性及活性氧代谢的影响[J].应用生态学报,2013,24(7):1809-1816.JING D W,XING S J,DU Z Y,et al.Effects of drought stress on the growth,photosynthetic characteristics,and active oxygen metabolism of poplar seedlings[J].Chinese Journal of Applied Ecology,2013,24(7):1809-1816.(in Chinese with English abstract)